Bone aerophones from Eynan-Mallaha (Israel) indicate imitation of raptor calls by the last hunter-gatherers in the Levant

Direct evidence for Palaeolithic sound-making instruments is relatively rare, with only a few examples recorded from Upper Palaeolithic contexts, particularly in European cultures. However, theoretical considerations suggest that such artefacts have existed elsewhere in the world. Nevertheless, evidence for sound production is tenuous in the prehistoric archaeological record of the Levant, the study of music and its evolution being sparsely explored. Here we report new evidence for Palaeolithic sound-making instruments from the Levant with the discovery of seven aerophones made of perforated bird bones in the Final Natufian site of Eynan-Mallaha, Northern Israel. Through technological, use-wear, taphonomic, experimental and acoustical analyses, we demonstrate that these objects were intentionally manufactured more than 12,000 years ago to produce a range of sounds similar to raptor calls and whose purposes could be at the crossroads of communication, attracting hunting prey and music-making. Although similar aerophones are documented in later archaeological cultures, such artificial bird sounds were yet to be reported from Palaeolithic context. Therefore, the discovery from Eynan-Mallaha contributes new evidence for a distinctive sound-making instrument in the Palaeolithic. Through a combined multidisciplinary approach, our study provides important new data regarding the antiquity and development of the variety of sound-making instruments in the Palaeolithic at large and particularly at the dawn of the Neolithic in the Levant.

www.nature.com/scientificreports/ Sound production in the Natufian of the Levant has previously been suggested, with only a few studies hinting at possible media/instruments for such a practice: a "belt" of bone pendants interpreted as strung rattles 25,26 , fragments of worked bone objects interpreted as bullroarers 26,27 , a fragment of a worked vulture ulna interpreted as a flute without holes 27 or even the supposed sound of pounding boulder mortars involved in funerary practices of Late Natufian contexts 28 . Altogether, sound-making instruments in the prehistoric Levantine archaeological record are yet to be thoroughly explored, indicating the great potential for further research. Thus, the multidisciplinary methodology developed in this article serves as the first systematic attempt to approach the issue. The aerophones from Eynan-Mallaha are all made of wing long-bones (one humerus, five ulnae, one radius) whose diaphysis have been perforated one to four times to form finger-holes. In the three cases where the epiphysis is still present, it has also been perforated to form the mouthpiece or the distal end of the object. To these finger-holes are added markings on three bones, either notches ( Fig. 2.7) or a series of small parallel incisions located near the finger-holes ( Fig. 4.2, 4.6), which are potentially linked to the placements of the fingers on the instrument. All the worked areas show contact-wear traces indicating that all instruments have been used. When looking at the state of preservation, most of the fractures are old, but some were caused by the excavation process. It is the case of the complete aerophone ( Fig. 2.7), which was broken in three pieces when discovered in 1998 and glued shortly thereafter, as were several other broken bird bones. Most of the remains are unburnt bones, although the shade of the humerus aerophone ( Fig. 2.6) indicates that it may represent a heated bone. As several bones and their technical traces are partially covered by encrustation, the micro-CT-Scan helped us to overcome the difficulty of analysing the bone surfaces (Fig. 3). Attributing the perforated bones from Eynan-Mallaha to acoustic instruments is based on the criteria developed by F. d'Errico and G. Lawson 29 , among them: (1) feasibility, (2) ethnographical parallels, (3) ancient documentary support (text or image), (4) contemporary archaeological support, and (5) efficiency. It is also based on an extensive archaeological and ethnographical corpus of such devices from diverse periods and regions 22,24,[30][31][32][33][34] .

Results
The makers of the aerophones carved the instruments from bird bones identified as the Eurasian teal (Anas crecca) and the Eurasian coot (Fulica atra). Both species are represented equally in the avifauna assemblage (respectively 11.2% and 10.7% of the Number of Identified Specimens: NISP) 5 (T.S., in prep). Four aerophones have been identified only at the level of Anas sp. (surface-feeding ducks) and are in the size range of several small ducks identified at the site (e.g., Garganey: A. querquedula, Pintail: A. acuta, Shoveler: Spatula clypeata). These small ducks are all winter migrants to the Levant and together represent 17.3% (n = 193) of the total NISP. The preferential selection of bird wing bones with their empty diaphysis, offering a long air pipe, for making aerophones corresponds to a universal trend in sound-making instruments as seen in the archaeological and ethnographical record 22,23,30,35 . Nevertheless, the choice to use the bones of small birds raises questions. Indeed, the Natufians from Eynan-Mallaha hunted other, larger species, such as birds of prey (Accipitridae), larger waterfowl (Goose, Swans) and especially the Mallard (Anas platyrhynchos), which alone represents 22.5% (n = 250) of the total NISP. Thus, selecting short and narrow bird bones as blanks for wind instruments appears to be more of a deliberate choice rather than a constraint of availability. Given that the length and diameter of the bird bone influence the sound production 36 it seems that the choice is less about the species used than the sound produced by these bones whose air pipe is smaller. This choice is not without consequence, as experiments have shown that the narrower the diameter of the bone is, the more difficult it is to play 36,37 .
These parameters also indicate that, contrary to what is observed in different archaeological cultures of the European Upper Palaeolithic (ulna in Isturitz and radius in Geissenklösterle 36 ), the fact that the Eynan-Mallaha Natufians used different wing bones (humerus, ulna and radius) to make aerophones probably reveals the search for varied sound production. This assumption will be tested with further experimental replicas made with humerus and radius bones. The small size of these instruments raises other questions. Indeed, if one considers that their size, in their functional state, corresponds at most to that of the complete aerophone (length of 63.4 mm and a diameter of about 4 mm (Fig. 2.7)), the mastery of these instruments must have required a certain period of training and a level of dexterity. The short distances between the finger-holes of certain aerophones ( Fig. 2.1;2.5; Figs. S1; S5; S9) (e.g., 2 mm from 1a to 1b or 5a to 5b) also requires a certain agility. Training with www.nature.com/scientificreports/ the experimental replicas and comparison with similar-sized bone aerophones from ethnographic contexts 38 will allow us to expand on this point.   Table S2), the extremities of the bones have been perforated by pressure ( Fig. S11; Fig. S12; Fig. S13) and bear signs of contact use-wear of varying intensity. On the complete ulna aerophone (Fig. 2.7), the proximal epiphysis has been perforated twice ( Fig. 2.7 g; Fig. S10) (caudal and cranial views) to form the mouthpiece. Those two perforations are not equal in size (4.8 mm 2 and 1.9 mm 2 ), which might have been done on purpose to influence the airflow, given that the mouthpiece shape conditions the sonority and the way of playing the instrument 22 . The mouthpiece displays a fairly well-developed contact use-wear since it extends further on the diaphysis. According to the typology of aerophones 22 , the mouthpiece allows us to classify the complete aerophone of Eynan-Mallaha as a flute with an unfabricated air duct because the player makes the duct during play with his/her mouth. As it has been demonstrated with the experimental replicas (Audio S1; Fig. S14), the air flow is brought in front of the edge of the perforation ( Fig. 2.7g 1 ; Fig. S10) which forms the bottom of a notch. Thus, we can classify the Eynan-Mallaha aerophone as a notched flute (like the traditional Quena flute from the Andes), one of the most difficult to play because the player has to hold the instrument in place against his/her lips so that the breath reaches the precise operational point (Fig. S14). According to Hornbostel and Sachs' classification of musical instruments 39 , it could be related to the category of "edge instruments or flutes (narrow stream of air is directed against an edge)".
On the distal end of the complete instrument, there is a single perforation ( Fig. 2.7j) (cranial view), which forms the end of the air column and whose size (3.05 mm 2 ) is probably not random. The only other distal end perforation preserved in its original shape ( Fig. 2.5; Fig. S9.5c), made on a distal ulna, is twice as large (6.4 mm 2 ). Following the same trend, the distance between the last diaphysis perforation and the distal end perforation is shorter on the complete instrument ( Fig. 2.7e-j = 11 mm) than on the broken one (  Table S2), were meticulously made with a short cutting edge (flint burin-bit angle) by transverse and oblique micro-grooving (Fig. 4). The grooving is precise and was obtained by movements alternatively from the right and left edges, reorienting the bone blank several times. The resulting finger-hole margin is in the form of a slightly concave plateau to the fingertip ( Fig. 4; Fig. S1-7). This shape serves to improve the pneumatic efficiency of the fingertip seal, which is acoustically essential for the wind instrument to produce the desired output sound tone 30 . The perforation by micro-grooving technique is never used elsewhere in the production of bone tools and ornaments in Eynan-Mallaha, where perforation by rotation is preferred [40][41][42][43] . The choice of this unusual technique could result from the physical and technical constraints posed by the small size and fragility of the bird bones used to make the aerophones. Indeed, with small size convex areas to be worked and a compact bone thickness of about 0.5 mm, it might have been challenging to control a perforation by rotation. This hypothesis is confirmed by the experimental replicas and by the aerophone made on a humerus ( Fig. 2.6), of which one of the perforations (Fig. S9.6a) was started by grooving and then finished by scraping in rotation, probably owing to a greater thickness of compact bone than on the other wing bones used. In the case of the largest finger-hole  Table S2). The finger-holes can be oval and irregular (Fig. 2.1 and 2.2) to circular and regular (Fig. 2.5 and 2.7). This difference in shape is accompanied by a difference in size, the perforations ranging from 0.35 mm 2 (Fig. S9.6a; Fig. 2.7i) to more than 3 mm 2 (Fig. S9.1b = 3.1 mm 2 ; Fig. 2.7c = 3.3 mm 2 ), allowing a ten times bigger air flow. As suggested by Zhang 34 for Neolithic Chinese bird bone flutes, the smallest perforations could have been made to correct the off-pitch tone of the bigger finger-holes. On the complete aerophone the distances between the finger-holes are varied, but a pattern is recognisable. There are two different spacings with a large finger-hole ( Fig. 2.7c) separated, by a large empty space (11.1 mm), from a concentration of 3 smaller finger-holes ( Fig. 2.7d,e,i). On www.nature.com/scientificreports/ three fragmented instruments made on ulna of a similar size ( Fig. 2.1, 2.2 and 2.5), we note a regularity in the spacing of the perforations ranging from 2 to 2.8 mm (Table S2), suggesting that the position of the finger-holes may have been predetermined, possibly by use of a template. As the bone size and the number and spacing of the finger-holes influence the sonic output, notably the frequency spectrum 36 , the similarity between them may suggest an attempt to achieve a similar range of sounds by standardising the instruments. On three of the instruments, markings ( Fig. 2.2;2.6;2.7; Fig. S9), either shallow notches made by transverse and oblique grooving (Fig. 2.7) or an oval concentration of small parallel incisions (Fig. 4), can be observed on the side of one of the finger-holes. Most probably, those markings were linked to the placement of the fingers on the instruments. On the complete aerophone, three additional markings, in the form of shallow notches ( Fig. 2.7a,b,f), are not related to any finger-holes and might be associated with the movement of the fingers during a sequence of play 24,30 .
The technical traces described leave no doubt that the holes are a product of intentional human activity. Indeed, they are very different in size, shape, and distribution from taphonomic alterations (e.g., carnivore puncture, digestion, rodent gnawing (Fig. 4), roots imprints, insect perforations, etc.) that we can observe on other bird bones in Eynan-Mallaha or even elsewhere within the research literature (see references in the Methods section). Moreover, the Natufian technical choices are similar to those made for the Aurignacian and Gravettian aerophones of Geissenklösterle and Isturitz caves 30,36 as well as several later examples in Europe 22 , the Americas 31-33 and China 34 . It is notably the case of the micro-grooving technique used to perforate the finger-holes, their margin in the form of a slightly concave plateau to the fingertip, incised markings near the finger-holes, the choice to perforate the convex side of the bone as well as the pattern of two different spacings with a large finger-hole separated, by a large empty space, from a concentration of three smaller finger-holes.
The use-wear on the outer finger-hole margin (Fig. 4) suggests that all the instruments have been used but at different intensities. Although more experimental references are needed, some use-wear studies on Palaeolithic aerophones 30,44 seem to demonstrate such assumptions. The only humerus aerophone (Fig. 4.6) showed the least intense wear, while one of the ulna aerophones ( Fig. 4.2) exhibited the most intense wear. None of the aerophones from Eynan-Mallaha show incisions commonly interpreted as decorations 22 , but residues of a red colouring matter can be seen on the shaft and mouthpiece of the complete aerophone ( Fig. 2.7g 3 ). These residues have been characterised as a mix of clay and ochre (hematite) by Scanning Electron Microscopy with Energy Dispersive X-ray Spectroscopy (SEM-EDS) (Fig. S10). They resemble the hundreds of others that we have observed on the Natufian shell and bone beads from Eynan-Mallaha 40 and, in the corpus of bird remains, this is the only occurrence of a bone-bearing residues of colouring matter, disproving a possible taphonomic origin of the residues. The presence of ochre per se in the archaeological remains and the presence of modified ochre fragments have been considered as part of the symbolic behaviour of prehistoric societies of diverse ages and geographical origins [45][46][47][48] . Moreover, the association of red colouring or red patterns with aerophones is also documented in the prehistoric 49 and ethnographic records 50 .
We reproduced experimentally three aerophones similar to the complete one of Eynan-Mallaha (Fig. 2.7). Replicas were made on both green and dry ulna of two female mallard individuals (Anas platyrhynchos) (Fig. S11). We used this species, close in size and shape (for female individuals), because of the difficulty in obtaining carcasses of Eurasian coot (Fulica atra) used by the Natufians. Our main goals in reproducing the marks observed on the archaeological objects were to assess the technical gestures involved in their production and to demonstrate the possibility of producing melodic (e.g., diverse tonalities and frequencies) sounds with the aerophone. The experimental marks are entirely consistent with those observed on the archaeological aerophones from Eynan-Mallaha. We showed that the Natufian artisans proceeded by pressure on the epiphysis and transversal grooving plus rotational scraping to perforate the finger-holes ( Fig. S11; Fig. S12; Fig. S13). We also reproduced several high-quality and high-pitched notes (maximum intensity approximately 65 dBA at 1 m of the instrument) on frequencies and levels in the human auditory field ranging from 2500 to 12,000 Hz ( Fig. 5A; Audio S1). We hypothesize that the purposes of the sound produced were to imitate bird calls. In that case, we can see that the three intense high frequencies produced by the replica (3000-4200 Hz, 4400-5600 Hz and 6050-7650 Hz) are not identified in the calls of Anas crecca (Fig. 5B) and Fulica atra (Fig. 5D) (Fig. S15), the species of which Natufians chose to use the bones to make the aerophones ( Table 2). Among the 58 species identified in the Final Natufian of Eynan-Mallaha 7 (Table S1), the calls of the Common kestrel (Falco tinnunculus) and the Sparrowhawk (Accipiter nisus) are the only ones that develop similar sound spectra ( Fig. 5C; Fig. S16). These two Falconiformes are the most familiar and easily seen raptor. According to the sonic affinity theory 51 , which holds that the action of surrounding sounds shapes the human musical brain, the calls of those familiar raptors should have therefore integrated the acoustic brain of the Natufians. Even if the Common kestrel and the Sparrowhawk have not been discovered in large numbers in Eynan-Mallaha (MNI = 3), they are represented almost exclusively by their talons (NISP = 10 out of 13), some of them bearing anthropic modification traces (LD, in prep) (Fig. 6), which indicates their use as personal ornaments and their symbolic status for the Natufians [19][20][21] . It should also be noted that these two species alone represent the majority (NISP = 10 out of 17) of the raptor talons found in the Final Natufian layer (17 species in total).
We, therefore, believe that the Eynan-Mallaha aerophones were made to reproduce the calls of the valued Common kestrel and Sparrowhawk. This function explains the Natufians' specific choice to use small blanks with reduced air conduct in order to create high-pitched sounds similar to the bird calls. But if the aerophones were functioning as bird calls, what was their purpose? They could have been meant as a decoy used to lure the Common kestrel and Sparrowhawk to facilitate their hunting (i.e., luring birds within shooting distance by imitating their sounds 52,53 ) but, with a total of seven aerophones made for only three of those birds identified on the site, the decoy would have lacked effectiveness. In contrast, ethnographic and archaeological evidence from various parts of the world consistently show that in societies where birds by-products (talons, feathers) are used as personal ornaments, vocal and instrumentally made bird imitative sounds have high symbolic value www.nature.com/scientificreports/ in traditional music and dance 50,54 . For example, during the Sun Dance of the tribes of Plains Indians, a major communal religious ceremony that reflects their relationships with nature, dancers imitate constantly and in unison the call of the symbolically valued Spotted eagle (Clanga sp.) by blowing in a whistle made of an eagle ulna bone decorated with feathers and red dots of ochre 50 . For the Kaluli (Great Plateau of Papua New Guinea), rainforest birds are considered to be spirits and their feathers are worn as ornaments. During women's funerary song weeping and men's ceremonial poetic songs, bird imitative sounds are intimately associated with the vocal music which allows the bird spirits to speak with the community through the musician 54 . We, therefore, wonder if imitative bird calls were integrated into Natufian musical or dancing practices. All the aerophones were found discarded with many other forms of occupational wastes of layer Ib (see 14 for detailed spatial distribution patterns and identified activity stations inside the shelters). The complete aerophone and a fragment also made from an ulna (Fig. 2.5 and 2.7) were found in two adjacent squares within 20 cm depth inside the roofed part of Shelter 200 (Fig. 6). The majority of the remains of Anas crecca and Fulica atra as the upper wing bones (humerus, ulna, radius) (Fig. 6) and the pectoral girdle bones (scapula, furculum, sternum and coracoid), associated with the largest meat mass in the avian body, are also concentrated in this structure (Fig. S17). There is another concentration of four instruments (Fig. 2.1,2,3,4) in two squares within 10 cm depth containing the Shelter 203 and the stony layer outside it (Fig. 6). This structure concentrates the upper wing bones of a bigger waterfowl, the Mallard (Anas platyrhynchos) (Fig. S17), which are nearly absent of the roofed part of Shelter 200 and have not been chosen to make aerophones. The thin archaeological horizon between these aerophones, inside the two areas of concentration (Shelter 200 and outside of Shelter 203), suggests a possible contextual link between them. Shelter 200 and Shelter 203 are also the contexts where most of the talons of the Common kestrel, the Sparrowhawk (NISP = 7 out of 10) (Fig. 6) and the other diurnal raptor have been found.
These aerophone concentrations suggest that there may have been areas of specialised activity, either for storing, playing or discarding sound-making instruments. This possibility is well illustrated by the complete aerophone discovered in the roofed part of Shelter 200 (square H91a), next to an activity station 14 where the upper wing bones of the birds chosen to make aerophones seem to have been curated (Fig. 6). The only humerus aerophone (Fig. 2.6) was found isolated, deeper in the slope of the stony layer, east of Shelter 215/228, where many bird bones have been discarded (Fig. 6; Figs. S17-S18). www.nature.com/scientificreports/

Discussion
The seven aerophones from the Final Natufian of Eynan-Mallaha share a specialised "chaîne opératoire" characterised by a unique manufacturing technique. Given the discretion of the technical traces on the bones, one could expect that similar instruments, not yet identified as such, are still hidden in the avifauna collections of other Natufian sites. We must also remember that the ethnographic record suggests a vast range of sound-making instruments are made from perishable materials 23 . In any case, we can already note that the seven aerophones of Eynan-Mallaha form the largest assemblage of prehistoric sound-producing instruments in the Levant. In a www.nature.com/scientificreports/ broader perspective, the complete aerophone of Eynan-Mallaha is one of the very few prehistoric sound-making instruments that has come to us in a complete state. Following musicological terms, music is sounds and notes in their social context, developed during social activities 55 . While the acoustic action is universal (all existing human societies produce vocal or instrumental music), the form and the means it takes are contingent and varied 22 . From the archaeological perspective, one fundamental problem is distinguishing between an artefact's potential to produce sound and the confirmation that this artefact served that specific purpose 29,55,56 . Hence, the global Palaeolithic record of sound-making instruments is scarce too. The earliest secured examples are aerophones from early Aurignacian contexts in southwestern Germany 24,30,36,57 . These examples fit into the archaeological paradigm of modern behaviour and the biological diversity of hominins in the Palaeolithic. Musical instruments are also recognised in later Upper Palaeolithic corpora in Europe (Gravettian and Magdalenian), including, for example, various aerophones and idiophones 58,59 . However, the evolution of music still needs to be completed.
Theoretical considerations suggest that music evolved through time and space 23,55,[60][61][62][63][64][65] . Music, on a phenomenological level, articulates with various aspects of our being, including brain functioning, behaviour, communication and socio-cultural traits [66][67][68] . Thus, positive paleo-organology (i.e., the research of ancient sound-making instruments) suggests new venues for future research-the prehistory of sound manipulations within their socio-cultural contexts through time and place. This emergent paradigm may expand our knowledge of various functions of different sounds and their co-evolution with biological (brain), behavioural, and, importantly, socio-cultural transformations in our prehistoric pasts.
Our study of the seven Natufian aerophones show that these instruments, meticulously manufactured, demonstrate the existence of a distinct category of objects that might represent a tradition of sound production in Eynan-Mallaha. These wind instruments employ the musically effective finger-hole principle for generating and organising of different sound pitches-the single-player melodic capability 30 -resulting in complex soundcommunication behaviour.
Altogether, technological, use-wear, taphonomic, experimental and acoustical evidence combined with comparative record of ethnographic and archaeological examples suggest a new type of Palaeolithic aerophone, which was not identified before. It is unique in the Palaeolithic record of sound instruments both by the chosen medium-i.e., small-sized bones of waterfowl, as well as the sounds it produces which closely resemble falcon calls. This new discovery contributes to the larger picture of music evolution (music sensu-lato) a yet unexplored category of sound-making instruments which have been part of the Palaeolithic acoustic environment, and produced artificial sounds resembling natural ones.
Moreover, here we add crucial evidence for the acoustic phenomenon of sound manipulation from a cultural context, which marks a significant change in the history of humankind-the transition into complex agricultural societies manipulating their vegetal and animal environments and accelerating the emergence of new ways of life.
The Natufian's manipulations of sounds might have functioned in various aspects of their socio-cultural lifeways, either for hunting, communication or ritualised behaviour 25 . Further exploration of this discovery may develop such questions as-what is the nature of the sound produced? How varied or restricted are they? What can we learn about their perception? And, how do they affect human and animal behaviour? Future research into these specifications, concerning the function, perception and effects of the sounds produced by these bone aerophones will be one of the foci of a more in-depth analysis that we will develop in the future. It is now clear that the evolution of music at the transition to agriculture, which articulated the intensification of socio-cultural complexity, was more branched than we supposed before. Thus, the exploration of Natufian acoustics gives a new perspective on this crucial period in human history.

Materials and methods
Archaeological methods. All  Generally speaking, the excavation was conducted using a unit system of a quarter square meter (0.25 m 2 ), 5 cm deep. This system was adapted according to the findings. The undifferentiated top of Layer Ib was excavated using a 1 m 2 grid, shifting to the quarter square meter grid inside and outside structures as soon as they were identified. The fill of each structure was isolated, introducing 'natural' boundaries instead of the artificial limits established by the grid. Nevertheless, the artificial grid subdivisions were respected in each structure to allow spatial distribution of the small finds.
The thickness of the units was also adapted to the stratigraphy, resulting in units much thinner than 5 cm, especially when searching for 'floors' and surfaces. As a rule, only exceptional pieces were spatially recorded during excavation in the undifferentiated upper part of Layer Ib, which was crowded with small limestone blocks. Inside the structures, all items larger than 5 cm (bones, stones, lithic artefacts and other finds) were mapped. Sediments were wet-sieved using a 1-2 mm mesh and further sorted in the laboratory.
Taxonomical and anatomical identification. Each bird bone retrieved has been numbered individually by the catalogue numbers of the excavation. All variables regarding the definition of taxa, element, completeness, and surface modifications were recorded in a database. All the remains are now housed at the National Natural History Collections at the Hebrew University of Jerusalem, Israel (NNHC-HUJ). The taxonomic identification of the bones took place in several phases of research spanning some 20 years. Initial identifications of taxa were conducted using the University of Michigan Museum of Zoology's avian collection. Later work, from ca. 2003 to this publication, was performed at the National Natural History Collections at The Hebrew University of Jerusalem, Israel (NNHC-HUJ). www.nature.com/scientificreports/ Taphonomic analyses. Each bone was examined under a light microscope (magnification 10-40×) and a Dino-Lite AD-7013MZT digital microscope (magnification 30-250×) for surface modifications caused by abiotic and biotic processes, such as carnivores, raptors, rodents, roots and human predation [69][70][71] . We have distinguished the technical traces from others, characteristic of gnawing, but also perforations, tearing and depressions caused by insects or the beaks, claws, teeth and talons of scavengers or birds of prey [72][73][74][75][76][77][78][79][80][81] . Anthropic perforations were differentiated from natural holes based on five main criteria: (i) anatomical location on the bone; (ii) size and nature of the marks; (iii) mode of manufacture; (iv) shape and regularity of the holes; (v) experimental perforations comparison (see description above and Fig. S11; Fig. S12; Fig. S13). We recorded every element and factor (time, number of incisions, the position of the hand, etc.) in a specific file designed for this experiment. Moreover, we graphically recorded all the processes with a Canon 6D camera (180 mm, 60 mm and 35 mm Tamron Macro lenses). The experimental marks were also analyzed microscopically with a stereomicroscope Kiowa SDZ-TR-P completed with a microscope camera Motic Moticam-S6. We conducted spectral analyses of the sounds produced by the replica, and bird calls with the free computer softwares Praat (version 6.2.23) and Spek (version 0.8.2). The bird calls of the 58 species identified in Eynan-Mallaha were sampled from the Sonothèque du Muséum national d'Histoire Naturelle of Paris (https:// sonot heque. mnhn. fr/) and Xeno-canto.org powered by the Xeno-canto Foundation and Naturalis Biodiversity Center (https:// xeno-canto. org/).